Fl. China 19: 104–141. 2011.

28. GALIUM Linnaeus, Sp. Pl. 1: 105. 1753. 拉拉藤属 la la teng shu

Chen Tao (陈涛); Friedrich Ehrendorfer

Subshrubs to perennial or annual herbs. Stems often weak and clambering, often notably prickly or “sticky” (i.e., retrorsely aculeolate, “velcro-like”). Raphides present. Leaves opposite, mostly with leaflike stipules in whorls of 4, 6, or more, usually sessile or occasionally petiolate, without domatia, abaxial epidermis sometimes punctate- to striate-glandular, mostly with 1 main nerve, occasionally triplinerved or palmately veined; stipules interpetiolar and usually leaflike, sometimes reduced. Inflorescences mostly terminal and axillary (sometimes only axillary), thyrsoid to paniculiform or subcapitate, cymes several to many flowered or in-frequently reduced to 1 flower, pedunculate to sessile, bracteate or bracts reduced especially on higher order axes [or bracts some-times leaflike and involucral], bracteoles at pedicels lacking. Flowers mostly bisexual and monomorphic, hermaphroditic, sometimes unisexual, andromonoecious, occasionally polygamo-dioecious or dioecious, pedicellate to sessile, usually quite small. Calyx with limb nearly always reduced to absent; hypanthium portion fused with ovary. Corolla white, yellow, yellow-green, green, more rarely pink, red, dark red, or purple, rotate to occasionally campanulate or broadly funnelform; tube sometimes so reduced as to give appearance of free petals, glabrous inside; lobes (3 or)4(or occasionally 5), valvate in bud. Stamens (3 or)4(or occasionally 5), inserted on corolla tube near base, exserted; filaments developed to ± reduced; anthers dorsifixed. Inferior ovary 2-celled, ± didymous, ovoid, ellipsoid, or globose, smooth, papillose, tuberculate, or with hooked or rarely straight trichomes, 1 erect and axile ovule in each cell; stigmas 2-lobed, exserted. Fruit on pedicels sometimes elongating during development, green, gray, or infre-quently white (to red, orange, or black), mostly dry to leathery schizocarps, infrequently spongy, rarely ± fleshy and berrylike, ellip-soid to subglobose; schizocarps separating into 2 indehiscent mericarps, each with 1 seed, subglobose, ellipsoid-oblong, or reniform, smooth and glabrous to tuberculate and/or covered with trichomes often hooked and clinging; seeds small, grooved ventrally (i.e., adaxially); testa membranous; endosperm corneous; embryo curved; cotyledons leaflike; radicle terete, inferior.

More than 600 species: worldwide, mostly in meridional to temperate but also in alpine and arctic regions or in subtropical and tropical zones at higher elevations; 63 species (23 endemic, four of unconfirmed occurrence) in China.

Galium is by far the largest and most widespread genus within the tribe Rubieae (subfamily Rubioideae). According to the most recent contri-butions (Natali et al., Opera Bot. Belg. 7: 193–203. 1996; Ehrendorfer et al., Fl. Iranica 176: 1–287. 2005; Bremer & Eriksson, Int. J. Pl. Sci. 170: 766–793. 2009; Soza & Olmstead, Taxon 59: 755–771. 2010), this tribe is closest to Theligoneae, Putorieae, and Paederieae, and includes the following genera treated (or mentioned) in the present flora: Asperula, Cruciata Miller, Galium, Kelloggia, Leptunis, Microphysa, Phuopsis, Rubia, and Sherardia Linnaeus.

So far, the genera Cruciata and Sherardia have not been found in China yet but may be expected there because of their partly weedy character and widely adventive occurrence. They are included in the key below for future reference but not among the full generic presentations. Sherardia arvensis Linnaeus is widely distributed in warm temperate and high-elevation tropical regions and can be separated from Asperula, Phuopsis, Leptunis, or Galium by its terminal capitate inflorescences enclosed by leaflike bracts, its clearly developed calyx with 6 acute lobes, and its pink or violet corollas with well-developed funnelform tubes and 4 lobes.

Among the few Cruciata species, the W Eurasiatic C. pedemontana (Bellardi) Ehrendorfer appears occasionally as an adventive in warm temperate regions. It is common, e.g., in SE North America, and could be found in China too. Cruciata can be separated from Galium by its flowering stems with vegetative apices and the inflorescences consisting only of lateral axillary cymes on middle and lower stem nodes. These cymes are equal to or shorter than the subtending leaves when fully developed. In contrast, the inflorescences are mostly terminal and axillary and longer than the leaves in Galium.

The characters relevant for the taxonomy of Galium and other Rubieae deserve some comments. Life and growth forms are important, particularly with respect to the differentiation into half-shrubs, herbaceous perennials, and annuals. Stem and leaf posture, consistency, shape, and indumentum (e.g., pubescent or retrorsely aculeolate with recurved microhairs) are often quite diverse and may vary within species or even populations. The true leaves are always opposite and 2, but interpetiolar stipules may vary from inconspicuous and divided or simple to enlarged and leaflike, forming whorls of 4 or up to 6 and more. During seedling and shoot development all these taxa pass through the 2- and 4-whorl stage, but some taxa remain at this stage, while others continue to develop more numerous whorl elements toward the middle of their stems. This is a most informative differential character within Rubieae. Other relevant features relate to leaf shape, venation, texture, and particularly indumentum. Here, the presence of longer or shorter microhairs (use a lens) on surfaces and particularly margins as well as their forward or backward direction is of tax-onomic importance.

Other morphological characters decisive for Rubieae taxonomy concern the inflorescences (e.g., the position and structure of the cymes). Flower shape is essential for the traditional separation of the genera Asperula (with salverform, funnelform, or cup-shaped corollas) and Galium (with ± rotate corollas). It is now clear that there are transitions between these character states and that even closely related taxa may differ in this respect. So far, it has been possible to provisionally maintain Asperula and Galium by the transfer of obviously misplaced taxa and by using the presence or absence of bracts and bracteoles as a differential character for the two genera (see Ehrendorfer et al., loc. cit. 2005).

The indumentum of ovaries and fruit as well as fruit consistency also vary strongly within Rubieae. Informative are, for example, ± fleshy berries (as in Relbunium Bentham & J. D. Hooker, Rubia, and certain Galium taxa) vs. dry schizocarps or the presence vs. absence of hairiness and whether the trichomes are hooked (i.e., the fruit disperse as “stick-tights” on animals) vs. straight. However, the distinction between all these structures is arbitrary, and there are even transitions between trichomes and tuberculate protuberances of various shapes as well as between hairy and glabrous.

Fl. China 19: 104–141. 2011.

All this is well illustrated by Yang and Li (Bull. Natl. Mus. Nat. Sci., Taichung 11: f. 1. 1998). Furthermore, ovary and fruit indumentum and surface structures may change during development and sometimes vary genetically within species or even within populations, as in several Galium species. In general, authors in other regions have documented infraspecific variation from glabrous to densely hairy or tuberculate fruit but traditionally have only separated plants with hooked trichomes into different species. However, intrepid Chinese authors have easily combined these latter morphotypes, e.g., in G. dahuricum sensu W. C. Chen (in FRPS 71(2): 255. 1999), whereas Fl. Japan (3a: 238–239. 1993) distinguished G. manshuricum on the basis of this character. Only careful studies and field observations can clarify such cases, as in G. spurium, where the infraspecific variation of fruit, either smooth, tuberculate, or covered with hooked hairs, has been proven.

Further differential characters for the taxonomy of Rubieae come from the fields of palynology (e.g., number of colpi), karyology (e.g., deviations from the normal chromosome base number x = 11 in Asperula sect. Cynanchicae (Candolle) Boissier with x = 10 or in Galium sect. Aparinoides (Jordan) Grenier with x = 12; common occurrence of polyploidy), and reproductive biology. Most of the perennial Rubieae taxa have conspicuous hermaphroditic or andromonoecious flowers and inflorescences and are insect-pollinated and self-incompatible outbreeders (e.g., Phuopsis or G. boreale and G. verum). Nevertheless, for several annuals with small and inconspicuous flower aggregates selfing and autogamy have been documented (e.g., G. aparine, G. spurium, and Sherardia arvensis). Furthermore, polygamodioecy and dioecy occur in some groups (e.g., G. elegans). Up to now, only few and insufficient data from all these fields are available for Asian Rubieae species and have not been mentioned in FRPS. Nevertheless, such data are significant and will have to be addressed in more detailed future systematic Rubieae studies from this region.

The α-taxonomy of Rubieae in E Asia is still in a problematic state. A general survey of the collections at the herbaria KUN, MO, PE, W, and WU has revealed the existence of many very polymorphic, complex, and insufficiently understood species groups. Therefore, the present treatment has to be regarded as provisional.

A particularly critical case concerns several Galium species described by H. Léveillé from 1904–1917 (see Lauener & Ferguson, Notes Roy. Bot. Gard. Edinburgh 32: 103–115. 1973). These descriptions are most fragmentary and the relevant types are not yet studied sufficiently (but see Mill, Edinburgh J. Bot. 53: 193–213. 1996). Relevant taxa in alphabetical order are G. blinii (see under that name), G. bodinieri (see under G. blinii and G. rebae), G. cavaleriei (see under G. asperifolium), G. comarii (see under G. dahuricum), G. esquirolii (see under G. asperifolium), G. hongnoense (see under G. spurium), G. mairei (see under G. elegans), G. martini (see under G. bungei), G. quinatum (see under G. blinii), G. remotiflorum (see under G. bungei), and G. venosum (see under G. bungei).

The treatment of Galium for the Flora of Taiwan by Yang and Li (Bull. Natl. Mus. Nat. Sci., Taichung 11: 101–117. 1998; Fl. Taiwan, ed. 2, 4: 254–259. 1998) is not satisfactory in several aspects: keys and descriptions are rather idealized and lack carefully observed ranges of morphological variation for the taxa; species are circumscribed more narrowly and based on different characters than used by other authors in the region (e.g., presence vs. absence of leaf indumentum is considered variable within species by most other authors); the treatment is not well reconciled with continental Galium taxonomy (e.g., there are no references to the Russian floras, and names synonymized by others are used without explanation); and at least two names based on types from Taiwan are missing.

With respect to a more “natural” and general taxonomic classification of the Rubieae and Galium, a number of recent morphological, karyological, palynological, and particularly DNA-analytical studies (e.g., Natali et al., loc. cit.; Robbrecht & Manen, Syst. & Geogr. Pl. 76: 85–146. 2006; Bremer & Eriksson, loc. cit.; Soza & Olmstead, loc. cit.) are available. They show that Theligonum should be placed into a separate tribe (Theligoneae), that the tribe Rubieae is monophyletic, and that Kelloggia (as subtribe Kelloggiinae, still with normal Rubiaceae stipules, calyx teeth, and 3-colpate pollen, but already with hooked trichomes on the dry mericarps) occupies a basal position in Rubieae. The Central American genus Didymaea J. D. Hooker (still with normal stipules but with the calyx already lacking, 5-coplate pollen, and seeds separating from the fleshy pericarp) represents a link to the genus Rubia in the true Rubiinae. Their stipules are nearly always leaflike, the pollen is polycolpate, and the seeds never separate from the pericarp. Rubia, a well-circumscribed and certainly monophyletic genus, is always perennial, has 5-lobed corollas, and berrylike fruit.

The remaining Rubiinae are also monophyletic as a whole, but their traditional genera Asperula, Bataprine Nieuwland, Callipeltis Steven, Crucianella Linnaeus, Cruciata, Galium, Leptunis, Mericarpaea Boissier, Microphysa, Phuopsis, Relbunium, Sherardia, Valantia Linnaeus, and Warburgina Eig are all essentially interdigitated. They are difficult to separate and can hardly be brought into concordance with available phylogenetic data. These advanced Rubiinae tend to develop more and more apomorphic character profiles, i.e., change from perennial to annual, increase in numbers of leaflike stipules from 4 to numerous, loss of bracts and prophylls in the inflorescences, reduction from 5-lobed to (3 or)4-lobed corollas, specialization of mericarps, etc. As shown by the most comprehensive phylogram available so far (Natali et al., loc. cit.: f. 2; Soza & Olmstead, loc. cit.: f. 1, 2) and new findings (unpubl.), these more apomorphic Rubiinae form a polytomy or a grade with seven parallel clades. The most basal clade (1) consists of the monotypic Galium sect. Cymogalia Pobedimova only. The following Sherardia clade (2) includes Crucianella, Phuopsis, Sherardia, and several sections of Asperula together with Leptunis. Separate clades are formed by G. sect. Depauperata Pobedimova (3), A. sect. Glabella Grisebach, including G. sect. Aparinoides (4), and A. sect. Asperula (5). The Cruciata clade (6) consists not only of the genera Cruciata and Valantia but also of all sections of Galium (including the traditional genera Bataprine, Microphysa, and Relbunium) that form whorls of 2 leaves and normally not more than 2 even-sized leaflike stipules. Finally, the G. sect. Galium clade (7) comprises this and various other sections of Galium, which regularly develop whorls of leaves and leaflike stipules with 5 to more elements.

From the above data and the fact that a number of major groups of Rubiinae have not been DNA-analyzed yet, it is obvious that it is still difficult and partly impossible to harmonize DNA-supported clades with the traditional genera and sections. Thus, extensive changes are expected for generic and sectional circumscriptions within Rubiinae in the future. Therefore, we refrain from taxonomic changes for the present flora, list taxa in alpha-betical order, and only supplement phylogenetic comments. Thus, the present treatment in principle follows FRPS (71(2): 216–286. 1999), mainly based on Pobedimova et al. (Fl. URSS 23: 287–381. 1958), but also considers Ehrendorfer et al. (loc. cit. 2005). In order to make comparison with available phylogenetic data and present infrageneric classification easier, relevant information is inserted as a “Taxonomic Conspectus” before the individual species descriptions. It was not until this volume was ready for the press that the need for the nomen novum, Galium glabriusculum, was discovered; therefore, this species alone is outside of the alphabetical order.

Fl. China 19: 104–141. 2011.

Here the key to species of Galium is extensively revised from that of FRPS. It includes all of the Chinese Galium species with full ranges of differential character variation. Furthermore, it keys out all other Rubieae genera which are easily confused with Galium and are documented or can be expected in China. Details on the genera Asperula, Leptunis, Microphysa, Phuopsis and Rubia can be found where they are listed in alphabetical order, references to Cruciata and Sherardia appear in the comments above.

Several species are keyed out more than once in the present key because they are circumscribed by combinations of characters rather than by unique features. Furthermore, many Galium species are markedly variable because of genetic differentiation (e.g., G. bungei, G. elegans) but also because of phenetic plasticity due to different environmental conditions. References to the number of leaves and leaflike stipules in whorls as well as leaf measurements refer to middle stem regions. Measurements of organs with hairy surfaces (e.g., leaves, fruit, and mericarps) here apply to the solid surface of the structure and do not include the trichomes. The terms “leaf whorl,” “ovary,” and “uncinate trichome” follow common usage in Galium. Infraspecific taxa are adopted from FRPS in order to facilitate future and more detailed work on this group and comparison with other floras. They are not included in the following main key but are subordinated under the relevant species in alphabetical order and keyed out there.

Taxonomic conspectus of the Rubieae (excluding Kelloggia and Rubia) In FRPS (71(2): 216–286. 1999) the taxa of Galium were placed in the following sections (designated here by capital letters): G. sect.

Depauperata (A), G. sect. Aparine (B), G. sect. Pseudaparine Lange (C), G. sect. Cymogalia (D), G. sect. Trachygalium (E), G. sect. Leptogalium Lange (F), G. sect. Platygalium (G), G. sect. Galium (H), G. sect. Leiogalium (I), G. sect. Trichocarpa (Pobedimova) Pobedimova (J), G. sect. Asperu-loides Pobedimova (K), and G. sect. Brachyantha (Boissier) Pobedimova (L). Some species of uncertain position were classified as dubious (M). Species accepted in the present treatment but lacking in FRPS are designated as (Z). For comparison, a current (e.g., Jelenevsky et al., Novosti Sist. Vyssh. Rast. 35: 174–187. 2003; Ehrendorfer et al., Fl. Iranica 176: 1–287. 2005), DNA-analytically supported (e.g., Natali et al., Opera Bot. Belg. 7: 193–203. 1996; Soza & Olmstead, Taxon 59: 755–771. 2010; Ehrendorfer, unpubl.) but still provisional taxonomic conspectus is presented below. It lists all species here accepted under their clades and sections. For the clades 1–6 one should compare the comments above, for the sections compare Ehrendorfer et al. (loc. cit.). The placement of Galium species into sections (or lack of placement) by FRPS is indicated by showing the relevant letters used above in parentheses after the species names.

Clade 1 Galium sect. Cymogalia Pobedimova s.s.

39. Galium paradoxum (D)

Clade 2 Phuopsis stylosa Sherardia arvensis Leptunis trichodes Asperula oppositifolia

Clade 3 Galium sect. Depauperata Pobedimova

16. Galium exile (A; incl. G. songaricum sensu FRPS)

Clade 4 Galium sect. Aparinoides (Jordan) Grenier

27. Galium karakulense (E) 25. Galium innocuum (as G. trifidum: E;

incl. “G. palustre”) Clade 5 Asperula sect. Asperula

Asperula orientalis

Clade 6 Galium sect. Platygalium W. D. J. Koch s.l.

9. Galium bungei (E; incl. G. martini: M)

45. Galium salwinense (E) 12. Galium crassifolium (A) 31. Galium linearifolium (E) 21. Galium hirtiflorum (Z) 20. Galium glandulosum (A) 18. Galium forrestii (D) 44. Galium rupifragum (Z) 35. Galium morii (D) 54. Galium tarokoense (A) 34. Galium minutissimum (M)

36. Galium nankotaizanum (M; incl. G. maborasense)

47. Galium serpylloides (A) 29. Galium kinuta (G) 24. Galium hupehense (M) 30. Galium kunmingense (Z) 17. Galium formosense (as G. kwanzan-

ense: M) 15. Galium elegans (D) 63. Galium yunnanense (M) 26. Galium kamtschaticum (D) 10. Galium chekiangense (as G. nakaii:

G) Microphysa elongata 40. Galium platygalium (K) 32. Galium maximoviczii (K) 7. Galium boreale s.l. (G) 60. Galium turkestanicum (M)

Clade 7 [Galium s.s.] Galium sect. Hylaea (Grisebach) Ehrendorfer s.l.

37. Galium odoratum (J) 4. Galium asperuloides (A) 22. Galium hoffmeisteri (as subsp. of G.

asperuloides: A) 14. Galium echinocarpum (A) 53. Galium takasagomontanum (M) 59. Galium triflorum (A) 58. Galium trifloriforme (Z)

Galium sect. Trachygalium K. Schumann s.l. 48. Galium sichuanense (Z) 13. Galium dahuricum (as “G. davuri-

cum”: F; incl. G. comarii, G. nie-werthii, G. pseudoasprellum)

56. Galium tokyoense (as var. of “G. davuricum”: F)

41. Galium prattii (M)

52. Galium taiwanense (M) 3. Galium asperifolium (I) 6. Galium blinii (as syn. of G. asperifolium

var.: I; incl. G. quinatum: M) 51. Galium sungpanense (A) 42. Galium pusillosetosum (A) 1. Galium acutum (M) 43. Galium rebae (Z) 33. Galium megacyttarion (Z) 5. Galium baldensiforme (A) 49. Galium glabriusculum (A) 28. Galium karataviense (as G. rivale s.l.:

K) 61. Galium uliginosum (F)

Galium sect. Leiogalium Ledebour 38. Galium paniculatum (M; incl. G. xin-

jiangense: J) Galium sect. Orientigalium Ehrendorfer

8. Galium bullatum (I) Galium sect. Galium

23. Galium humifusum (L) 62. Galium verum (H) 11. Galium consanguineum (as G. maj-

mechense: H) 46. Galium saurense (M)

Galium sect. Aparine W. D. J. Koch s.s. 50. Galium spurium (as G. aparine var.

tenerum: B) 2. Galium aparine (B)

Galium sect. Kolgyda Dumortier s.s. 57. Galium tricornutum (as G. tricorne: B)

Galium sect. Microgalium Grisebach 19. Galium ghilanicum (Z) 55. Galium tenuissimum (C)

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Key to species of Galium and to related genera of the Rubieae 1a. Interpetiolar stipules inconspicuous, multifid or fimbriate, not leaflike and not forming whorls with true

leaves; corolla funnelform, (4 or)5-lobed; ovary and dry mericarps with hooked trichomes ........................ Kelloggia (see p. 183) 1b. Interpetiolar stipules mostly leaflike and in whorls with true leaves, rarely reduced.

2a. Corolla lobes regularly 5; fruit fleshy, mericarps berrylike, 2(or 1, by non-development), often dispersed together ................................................................................................................................................. Rubia (see p. 305)

2b. Corolla lobes usually 4 (rarely 3); fruit dry or leathery, mericarps mostly 2, nearly always separating for dispersal.

3a. Leaves in middle stem region opposite, with stipules reduced or ± leaflike and in whorls of 4 but then always clearly smaller than true leaves.

4a. Corolla pink, funnelform, with well-developed tube longer than lobes; fruit smooth ........ Asperula oppositifolia (see p. 78) 4b. Corolla white, rotate, with tube shorter than lobes; fruit with uncinate trichomes.

5a. Perennial herbs; leaves 5–30 × 5–23 mm, obtuse to truncate at base, on petioles 1.5–10 mm; flowers 3–11 in cymes; corolla with 4 lobes ......................................................................... 39. G. paradoxum

5b. Annual herbs; leaves 2–12 × 1–4 mm, acute to cuneate at base, subsessile or on short petioles; flowers solitary at each node; corolla mostly with 3 lobes .................................................................................. 16. G. exile

3b. Leaves in middle stem region opposite and with very similar leaflike stipules in whorls of 4–16. 6a. Leaf apex rounded, obtuse, or ± blunt, never acute or with a hyaline mucro; leaves in whorls

of 4–6, linear to broadly oblong, 1-nerved, dried blackening; ripe mericarps globose, didymous and only with a short zone of contact, glabrous; corolla cup-shaped to slightly campanulate, 3- or 4-merous.

7a. Inflorescences with many-flowered cymes; corolla 4-lobed, 2.5–4 mm in diam.; leaves mostly 15–20 × 5–8 mm ....................................................................................................................................... 27. G. karakulense

7b. Inflorescences with 1–3(or 4)-flowered cymes; corolla mostly 3-lobed, 1–1.8 mm in diam.; leaves mostly 3–8 × 1–2 mm ................................................................................................................................... 25. G. innocuum

6b. Leaf apex mostly ± acute, often with a hyaline mucro; leaves in whorls of 4–16, sometimes broader and with 3–5 palmate nerves; ripe mericarps ovoid to subglobose, with a longer zone of contact and with diverse surface structures; corolla diverse, but often rotate and always 4-merous.

8a. Leaves and leaflike stipules in middle stem region never in whorls of more than 4 (if rarely in whorls of up to 6 then leaves with 3–5 palmate principal veins), from linear to broadly ovate.

9a. Stem apex vegetative, with few- to several-flowered lateral cymes only in leaf axils and shorter than or ± equal to subtending leaves, nodding in fruit .................................. Cruciata (see comments above)

9b. Stem apex usually floriferous, with terminal and axillary cymes, often longer than subtending leaves and mostly not nodding in fruit.

10a. Condensed plants of rocks or high elevations; stems usually less than 10 cm tall, glabrous or with spreading (but never retrorse) hairs; leaves mostly ± ovate, (1–)3–8(–20) × (0.8–)2–4(–10) mm, with 1–3 main veins; corolla rotate, often only 1.5–2 mm in diam.; mericarps with spreading (rarely appressed) hooked or ± straight trichomes.

11a. Mericarps with ± straight hairs, 2–2.5 mm in diam.; stems mostly pilose or hirtellous. 12a. Fruiting pedicels straight; Xizang ............................................................................................. 47. G. serpylloides 12b. Fruiting pedicels nodding; Taiwan ....................................................................................... 36. G. nankotaizanum

11b. Mericarps with weakly to strongly curved and uncinate trichomes; stems partly glabrous. 13a. Corolla ca. 3 mm in diam.; stems glabrous .................................................................. 53. G. takasagomontanum 13b. Corolla 1.2–2 mm in diam.; stems ± hairy or glabrous.

14a. Stems ± hairy. 15a. Leaves ovate to broadly lanceolate, acute, up to 3.5 mm wide; Yunnan ............................ 44. G. rupifragum 15b. Leaves broadly elliptic to obovate, obtuse and mucronate, up to 10 mm wide;

Taiwan .................................................................................................................................. 17. G. formosense 14b. Stems glabrous; Taiwan.

16a. Leaves very small, only 0.8–1 mm wide, with 1 main vein only; fruit hairs spreading .......................................................................................................................... 34. G. minutissimum

16b. Leaves wider, with 1 or 3 main veins; fruit hairs appressed. 17a. Leaves with 3 main veins; corolla only ca. 1.2 mm in diam. .................................................... 35. G. morii 17b. Leaves with 1 main vein; corolla ca. 2 mm in diam. ........................................................ 54. G. tarokoense

10b. Taller plants, usually of lower elevations with larger leaves (if plants ± condensed then stem hairs retrorsely curved or fruit hairs appressed but not hooked).

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18a. Leaves with 1 principal vein or 2 lateral veins only weakly visible and not extending past middle of blade.

19a. Open corollas funnelform, 2.5–3 mm, tube somewhat shorter than lobes; fruit with pericarp smooth to granular, becoming slightly inflated, enclosing both mericarps at dispersal ................................................................................ Microphysa elongata (see p. 216)

19b. Open corollas rotate, fused basal part much shorter than lobes; mericarps clearly separated.

20a. Stems ± strigose-hirsute, with hairs ± retrorse (but not retrorsely aculeolate); leaves ovate or elliptic to linear-lanceolate, broadest ± in middle, at lower side usually with glandlike spots; flowers unisexual, usually yellowish, ± greenish, or reddish; fruit normally with uncinate trichomes.

21a. Plants usually less than 15 cm tall, strongly branched from base; leaves often less than 8 mm, mostly glabrescent or glabrous, subleathery; inflorescences with few-flowered, bracteate cymes .......................................................................................... 20. G. glandulosum

21b. Plants usually more than 15 cm tall, little branched; leaves usually longer than 8 mm, hairy on both sides; inflorescences paniculate to corymbiform, little bracteate.

22a. Leaves linear-elliptic to narrowly lanceolate, mostly 8–17 × 1–2.5 mm, dried rather papery; inflorescence paniculate ....................................................................................... 21. G. hirtiflorum

22b. Leaves ovate-elliptic, mostly 8–12 × 3–5 mm, dried rather subleathery; inflorescence corymbiform ........................................................................................................................... 18. G. forrestii

20b. Stems glabrous or with indumentum, but not with retrorse hairs; leaves often broadest above middle and thinner, without glandlike spots; flowers usually bisexual.

23a. Leaves ± linear, often longer than 20 mm, in addition to 1 principal, with 2 weaker lateral veins; corolla 4–5 mm in diam.

24a. Leaves linear-spatulate, 1–4 mm wide; inflorescences loose, broadly paniculiform; ovaries and fruit glabrous and smooth ........................................................................... 31. G. linearifolium

24b. Leaves linear-lanceolate, 3–9 mm wide; inflorescences dense, elongate-paniculate; ovaries and fruit with sparse hooked trichomes or glabrous ....................................... 60. G. turkestanicum

23b. Leaves not linear, mostly shorter than 20 mm; corolla smaller. 25a. Leaves ovate, length/breadth index 2 or less, in addition to 1 principal, with 2 weaker

lateral veins; corolla larger; fruit with spreading hooked or straight hairs; Taiwan. 26a. Mericarps with straight trichomes; corolla 2–2.5 mm in diam.; stems pilose or

glabrescent .............................................................................................................. 36. G. nankotaizanum 26b. Mericarps with hooked trichomes; corolla ca. 3 mm in diam.; stems

glabrous ............................................................................................................ 53. G. takasagomontanum 25b. Leaves ovate-oblong to lanceolate, length/breadth index 2 or more, with

only 1 principal vein; corolla 1.5–2 mm in diam.; ovaries and fruit glabrous or with various indumentum.

27a. Leaves dried subleathery; fruit with appressed, ± curved (but not uncinate) hairs; Shanxi ................................................................................................................. 12. G. crassifolium

27b. Leaves dried papery; fruit with various indumentum. 28a. Plants ascending, weak, sparsely hairy or glabrous; inflorescence few flowered,

peduncles and pedicels very thin and elongated, latter mostly 4–8 mm; fruit with spreading uncinate trichomes ............................................................................. 45. G. salwinense

28b. Plant erect, more robust, indumentum diverse; inflorescences ± many flowered, peduncles and pedicels thicker and shorter, latter mostly 2–4 mm; fruit tuberculate, with appressed or spreading hooked trichomes, or more rarely smooth .................................................................................................................................. 9. G. bungei

18b. Leaves with 3–5 palmate principal veins, lateral veins well marked and extending for more than half of blade length.

29a. Corolla funnelform or cup-shaped, 2–5 mm in diam., with fused lower part ± as long as lobes; ovaries and fruit glabrous.

30a. Corolla cup-shaped or campanulate, 2–2.7 mm in diam.; cauline leaves broadly lanceolate, always in whorls of 4 .......................................................................................... 30. G. kunmingense

30b. Corolla funnelform or campanulate, 2.5–5 mm in diam.; middle stem leaves ovate to elliptic, in whorls of 4–6.

31a. Open corollas 3.5–5 mm in diam.; cauline leaves usually in whorls of 4, 12–28 mm ............................................................................................................................ 40. G. platygalium

Fl. China 19: 104–141. 2011.

31b. Open corollas 2.5–3.5 mm in diam.; cauline leaves in whorls of 4–6(–8), 23–53 mm .......................................................................................................................... 32. G. maximoviczii

29b. Corolla rotate, (1–)2–5 mm in diam., with fused base much shorter than lobes. 32a. Leaves linear to linear-lanceolate, 27–40 × 3–9 mm, in addition to 1 principal, with

2 weaker lateral veins; corolla 4–5 mm in diam.; ovaries and fruit with sparse uncinate trichomes or glabrous ........................................................................................... 60. G. turkestanicum

32b. Leaves lanceolate to ovate, shorter, 3 principal veins mostly readily visible. 33a. Fruit glabrous, smooth to granular-papillose, or with appressed and ± hooked or with

spreading and straight (but never with spreading and hooked) trichomes. 34a. Open corollas 3–4 mm in diam.; stems (except nodes) mostly glabrous and smooth.

35a. Leaves ovate-lanceolate to ovate, papillose, length/breadth index mostly 2.5 or less; cymes rather few flowered; ovaries and fruit with ± appressed, apically somewhat bent trichomes .......................................................................................... 10. G. chekiangense

35b. Leaves mostly rather narrowly lanceolate, smooth or ± hairy, length/breadth index mostly 3 or more; cymes many flowered; ovaries and fruit glabrous or with various indumentum ...................................................................................................... 7. G. boreale

34b. Open corollas (1–)2–2.5 mm in diam.; stems glabrous or ± hairy. 36a. Cauline leaves broadly to narrowly lanceolate, length/breadth index often

3.5 or more. 37a. Stems hairy throughout; leaves lanceolate; fruit with straight hairs or rarely

glabrous ........................................................................................................................ 24. G. hupehense 37b. Stems (except nodes) glabrous; leaves ovate-lanceolate (sometimes also broader),

apex subacute to acuminate, striate-punctate glandular below; fruit glabrous and smooth .......................................................................................................................... 29. G. kinuta

36b. Cauline leaves narrowly to broadly ovate-lanceolate, length/breadth index usually 3 or less.

38a. Leaves 6–30 × 3–20 mm; fruiting pedicels straight; fruit glabrous or scaberulous; mainland ........................................................................................................................... 15. G. elegans

38b. Leaves 4–10 × 2–5 mm; fruiting pedicels nodding; fruit with ± straight trichomes; Taiwan ................................................................................................ 36. G. nankotaizanum

33b. Fruit with ± spreading and uncinate trichomes. 39a. Open corollas 3 mm or more in diam.; stems (except nodes) often glabrous and smooth.

40a. Leaves lanceolate to ovate-lanceolate or elliptic, length/breadth index 3.5 or more, apex acute to acuminate ......................................................................................................... 7. G. boreale

40b. Leaves ovate, length/breadth index less than 3. 41a. Leaf apex obtuse to rounded, usually mucronate; mainland ............................... 26. G. kamtschaticum 41b. Leaf apex acuminate; Taiwan ....................................................................... 53. G. takasagomontanum

39b. Open corollas 2.5 mm or less in diam.; stems often ± hairy. 42a. Leaves ovate-lanceolate to narrowly elliptic, with acute apex, length/breadth

index normally more than 2.5 ....................................................................................... 63. G. yunnanense 42b. Leaves ovate to broadly elliptic, with obtuse to rounded apex, entire or shortly

mucronate, length/breadth index normally less than 2.5. 43a. Leaves up to 20 mm wide; plants slender to usually rather robust; mainland ................ 15. G. elegans 43b. Leaves up to 10 mm wide; slender low plants; Taiwan ............................................ 17. G. formosense

8b. Leaves and leaflike stipules in middle stem region regularly in whorls of more than 4, i.e., in whorls of 5–16, with only 1 principal vein, linear to broadly lanceolate or elliptic, but never ovate or with a length/breadth index of less than 2.5.

44a. Inflorescences capitate and enclosed by leaflike bracts; corolla funnelform or salverform, 4–15 mm, with 4 or 5 lobes.

45a. Plants perennial, 20–70 cm tall; calyx limb obsolete; corolla 5-lobed, 12–14 mm ...... Phuopsis stylosa (see p. 291) 45b. Plants annual; corolla 4-lobed, shorter.

46a. Calyx teeth well developed; corolla pink, tube 4–5 mm ........................ Sherardia arvensis (see comments above) 46b. Calyx lacking; corolla bluish, tube 7–13(–15) mm .................................................. Asperula orientalis (see p. 78)

44b. Inflorescences branched, not enclosed by bracts; corolla rotate, campanulate, or funnelform, 0.5–13 mm, mostly with 4 (rarely 3) lobes.

47a. Medium stem leaves marginally (and often on upper side) with microhairs directed forward (use 20× lens), thus antrorsely ciliolate or aculeolate; stems mostly not retrorsely aculeolate.

48a. Ovaries and fruit densely covered with uncinate trichomes; plants perennial with smooth stems.

Fl. China 19: 104–141. 2011.

49a. Corolla funnelform, with tube ± as long as lobes ......................................................................... 37. G. odoratum 49b. Corolla rotate, with tube markedly shorter than lobes.

50a. Robust, procumbent to ascending plants often longer than 50 cm; inflorescences terminal and lateral with cymes in upper 2–4 nodes ................................................................................ 59. G. triflorum

50b. Slender erect plants, less than 50 cm tall; inflorescences predominantly terminal. 51a. Leaves 6–25 × 2–7 mm; open corollas ca. 2 mm in diam.; Taiwan (cf. also

53. G. takasagomontanum) .............................................................................................. 14. G. echinocarpum 51b. Leaves often larger; open corollas 1.5–3.5 mm in diam.; mainland.

52a. Leaves mostly in whorls of 7 or 8, (ob)lanceolate, length/breadth index mostly 3.5–4.5, subpetiolate; fruit with uncinate trichomes 0.6–0.8 mm; expected in Xizang .......................................................................................................................... 4. G. asperuloides

52b. Leaves mostly in whorls of 6, narrowly obovate to broadly oblanceolate, length/breadth index mostly 2.5–3.5, clearly petiolate; fruit with uncinate trichomes 0.8–1.2 mm or longer; widespread ................................................................. 22. G. hoffmeisteri

48b. Ovaries and fruit glabrous or hairy, but never with uncinate trichomes; plants perennial or annual.

53a. Plants annual, slender; flowers on pedicels and peduncles often longer than 5 mm and in lax, diffuse inflorescences.

54a. Leaves filiform, 20–30 mm, ascending; corolla funnelform, pink to red; ovary and mericarps with dense, short and curved hairs ............................................. Leptunis trichodes (see p. 213)

54b. Leaves linear to oblanceolate, 4–20 mm, spreading to reflexed; corolla ± rotate, whitish, yellowish, or greenish; ovary and fruit glabrous or slightly tuberculate.

55a. Inflorescences broadly ovate, diffuse and intricate, with fruiting pedicels elongated to 20 mm ............................................................................................................................. 55. G. tenuissimum

55b. Inflorescences rather narrowly thyrsoid, not diffuse and intricate, with fruiting pedicels only up to 4 mm ..................................................................................................... 19. G. ghilanicum

53b. Plants perennial, slender to robust; flowers on pedicels and peduncles 0.5–5 mm, in lax to ± congested inflorescences.

56a. Corolla funnelform, lobed for 1/2–2/3, white. 57a. Inflorescences lax, ± ebracteate; stems erect, smooth; main stem leaves 15–65 ×

3–12 mm ............................................................................................................................. 38. G. paniculatum 57b. Inflorescences congested, strongly bracteate; stems procumbent, mostly ± hairy;

main stem leaves 5–23 × 1–2(–5) mm ................................................................................. 23. G. humifusum 56b. Corolla rotate, lobed for 3/4 or more, often yellowish; plants erect to ascending.

58a. Leaves in middle stem region in whorls of not more than 6; plants of (sub)alpine region, not taller than 30 cm ................................................................................................................. 46. G. saurense

58b. Leaves in middle stem region in whorls of more than 6 and up to 12. 59a. Open corollas 3.5–5 mm in diam., white; fruit somewhat spongy or fleshy, 3–3.5 mm,

with a dry pericarp separating from rest of fruit ..................................................................... 8. G. bullatum 59b. Open corollas ca. 3 mm in diam., yellow to whitish; fruit with dry mericarps,

1.5–2 mm, with pericarp dark and firmly attached to rest of fruit. 60a. Leaves mostly 2.5–5 mm wide, glabrous abaxially; fruit ca. 1.5 mm ................... 11. G. consanguineum 60b. Leaves 1–2.5 mm wide, glabrous to densely pubescent abaxially; fruit 1.5–2 mm ............. 62. G. verum

47b. Medium stem leaves marginally (not on leaf surface) with microhairs directed backward (use 20× lens), thus retrorsely aculeolate or completely glabrous and smooth; stems often retrorsely aculeolate.

61a. Annuals, often in ± disturbed, weedy habitats; stems and leaf margins retrorsely aculeolate; fully developed mericarps subspherical, 2–6 mm; open corollas 1–2 mm in diam.

62a. Fruit becoming pendulous on arching peduncles and pedicels, verrucose to spinulose; leaves glabrescent above ............................................................................................................ 57. G. tricornutum

62b. Fruit on divaricate straight peduncles and pedicels (only latter sometimes bent just beneath fruit), mostly with uncinate trichomes; leaves ± hairy above.

63a. Open corollas 1.5–2 mm in diam.; individual mature mericarps 2.5–5 mm in diam., with trichomes arising from tuberculate bases ....................................................................................... 2. G. aparine

63b. Open corollas 1–1.5 mm in diam.; individual mature mericarps 1–3 mm in diam., with trichomes straight from base ......................................................................................................... 50. G. spurium

61b. Perennials, in ± natural habitats; fully developed mericarps ellipsoid, 1.5–3 mm; open corollas 1–4 mm in diam.

Fl. China 19: 104–141. 2011.

64a. Middle stem leaves narrowly obovate to oblanceolate, mostly 18–28 × 5–10 mm; stems slightly retrorsely aculeolate; inflorescences of medium size, with terminal and lateral, few- to several-flowered, rather loose cymes with small bracts, in fruit stiffly divaricate; corolla rotate, 1.5–2 mm in diam.; fruit with hooked trichomes .............................................. 58. G. trifloriforme

64b. Middle stem leaves mostly smaller; inflorescences different, usually more bracteate; fruit smooth, verrucose, or with hooked trichomes.

65a. Corolla funnelform to subcampanulate, with tube ± as long as or slightly shorter than lobes, whitish; ovaries and fruit glabrous, smooth or verrucose; leaves papery to subleathery and glossy; stems rough, retrorsely aculeolate, procumbent to clambering.

66a. Corolla funnelform; plants robust, 0.6–1.2 m, often forming mats; main stem leaves 8–50 × 2–8 mm .................................................................................................................. 28. G. karataviense

66b. Corolla subcampanulate; plants slender and ± erect, 10–60 cm tall; main stem leaves 3–16 × 1–3 mm ..................................................................................................................... 61. G. uliginosum

65b. Corolla rotate, fused basal part much shorter than lobes; ovaries and fruit with hooked trichomes, tuberculate, or glabrous; stems glabrous, rough, or hairy.

67a. Middle stem leaves larger, (5–)10–35(–50) × (1–)2.5–10 mm, mostly ± hairy, at least margins retrorsely aculeolate; cymes lateral and terminal, many flowered; ovaries and fruit glabrous or with diverse indumentum; plants from lower elevations, relatively robust, erect or clambering, stems up to 0.7 m tall.

68a. Stems branched from base; cymes leafy, with bracts to last branches; stems and leaves (nodes and margins excepted) glabrous and smooth; mericarps with spreading uncinate trichomes .......................................................................................... 48. G. sichuanense

68b. Stems normally branched from middle; cymes usually less leafy; stems and leaves mostly with more indumentum; fruit glabrous, papillose, tuberculate, or with uncinate trichomes.

69a. Peduncles and pedicels slender, filiform and often ± flexuose, with inconspicuous bracts; pedicels up to 5 mm, in fruit elongated to 10 mm or more; flowers never reddish; plants usually clambering; ovary and fruit surfaces diverse.

70a. Fruit often with uncinate trichomes; mainland .......................................................... 13. G. dahuricum 70b. Fruit glabrous; Taiwan ................................................................................................ 52. G. taiwanense

69b. Peduncles and pedicels rather stiff and often divaricate and ± bracteate; pedicels shorter than 5 mm and hardly elongated in fruit; ovary and fruit surfaces diverse, but often glabrous.

71a. Corolla red to purple (very rarely maroon or white), 1.5–2.5 mm in diam. ......................... 6. G. blinii 71b. Corolla whitish, yellowish, or greenish.

72a. Corolla small, 1.5–2 mm in diam., yellow to greenish white, lobes aristate; inflorescences divaricate and regularly bracteate with bracts similar to but smaller than leaves, giving a diffuse miniature aspect; ovary and fruit surface variable; plants often clambering ........................................................................... 3. G. asperifolium

72b. Corolla larger, mostly more than 2 mm in diam.; inflorescences ebracteate or with bracts ± reduced and irregularly distributed; ovary and fruit surface smooth to tuberculate; plants erect, hardly clambering.

73a. Leaves lanceolate, gradually narrowed into acute apex .............................................. 41. G. prattii 73b. Leaves subspatulate to obovate, apex rounded and abruptly narrowed into

a mucro ................................................................................................................... 56. G. tokyoense 67b. Middle stem leaves uniformly small, 2–12(–15) × 0.3–3.5 mm, glabrous and smooth

to ± hairy; cymes lateral and terminal, few flowered; ovaries and fruit with uncinate (very rarely ± straight) trichomes or glabrous; plants from high elevations, usually reduced and weak, caespitose to procumbent, stems only up to 0.3 m.

74a. Ovaries and fruit with uncinate (very rarely ± straight) trichomes. 75a. Leaves and stems ± densely hispid and often retrorsely aculeolate; stems with

4 conspicuous whitish angles .................................................................................. 42. G. pusillosetosum 75b. Leaves completely glabrous and smooth or only slightly hairy and/or retrorsely

aculeolate; stems with 4 inconspicuous angles. 76a. Leaves dried blackening, papery and thin, oblanceolate to narrowly obovate,

with flat margins, hardly longer than 7 mm ............................................................ 5. G. baldensiforme 76b. Leaves dried greenish-brownish, with ± revolute margins, often longer

than 7 mm. 77a. Plants nearly always smooth; leaves ± subleathery; ovary in flower

ca. 1 mm ............................................................................................................. 49. G. glabriusculum

Fl. China 19: 104–141. 2011.

77b. Plants retrorsely aculeolate at least on margins and lower side of papery leaves; ovary in flower 0.5–0.8 mm ................................................................................. 51. G. sungpanense

74b. Ovaries and fruit glabrous, smooth, papillose, or verrucose; Himalaya. 78a. Plants weak to procumbent but not mat-forming; cells of adaxial leaf surface

relatively large, readily visible individually with 20× lens; corolla mostly whitish ....................................................................................................................... 33. G. megacyttarion

78b. Plants procumbent and often mat-forming; cells of adaxial leaf surface small, not or hardly visible individually with 20× lens.

79a. Stems ± densely hairy and/or retrorsely aculeolate, with 4 conspicuous whitish angles .................................................................................................................... 42. G. pusillosetosum

79b. Stems glabrous or only slightly retrorsely aculeolate, with inconspicuous angles. 80a. Leaves on main stems 2–8.5 mm; inflorescence cymes 1- to few flowered,

fascicled; corolla white, pale green, or pale yellow, with upper surface of lobes papillose ................................................................................................................. 1. G. acutum

80b. Leaves on main stems 5–10.5 mm; inflorescence cymes 1–6-flowered; corolla nearly always red or purple, with upper surface of lobes glabrous and smooth except ± puberulent on margins and central vein ............................................................ 43. G. rebae

1. Galium acutum Edgeworth, Trans. Linn. Soc. London 20: 61. 1846.

尖瓣拉拉藤 jian ban la la teng

Herbs, perennial, procumbent, much branched, mat-forming. Stems up to 30 cm, 4(or 6)-angled, glabrous, smooth or sometimes with scattered (very rarely more dense) short and straight hairs. Leaves in whorls of up to 6, sessile; blade drying papery and blackish, linear-oblanceolate to narrowly elliptic-oblanceolate, 2–8.5 × 0.3–1.5 mm, glabrous and smooth, occa-sionally with straight hairs, base cuneate, margins flat to thinly revolute, very rarely antrorsely aculeolate, apex acute, ± con-tracted and mucronate; vein 1. Inflorescences with terminal and axillary cymes, 1- to few flowered; peduncles (1.5–)3–8(–10) mm; pedicels (0.1–)0.5–2(–3) mm, glabrous, smooth. Ovary ellipsoid-obovoid, ca. 0.5 mm, didymous, glabrous. Corolla white, pale greenish, or yellowish, rotate, 1.2–3.5 mm in diam., glabrous to puberulent, lobed for 2/3 or more; lobes 4, lanceo-late-spatulate, inside (i.e., adaxially) papillose, shortly acumi-nate. Mericarps ellipsoid, ca. 1 × 0.4–0.6 mm, glabrous, smooth or granular-verruculose, often on elongating pedicels. Fl. and fr. Jul–Oct.

Mountain rocks and slopes; 2000–4100 m. ?Sichuan, Xizang, ?Yunnan [India, Nepal, Pakistan].

Galium acutum is a (sub)alpine Himalayan member of the G. asperifolium group (see under that species). This group of taxa can be divided into subgroups: (1) from lower elevations and (2) from higher elevations. The latter subgroup is represented in the W Himalaya of Pakistan by G. acutum alone (Nazimuddin & Ehrendorfer, Pl. Syst. Evol. 155: 71–75. 1987). Mill (Edinburgh J. Bot. 53: 193–213. 1996; Fl. Bhutan 2(2): 825–834. 1999) has analyzed both subgroups in detail with emphasis on their E Himalayan members. Among subgroup (2) he recognized three species: G. acutum and the newly described G. rebae and G. megacyttarion. The only material from China incorporated in Mill’s study are specimens of G. rebae from Xizang deposited at E and BM. Additional species from subgroup (2) from the C and E Himalaya and adjacent China treated here are G. baldensiforme, G. pusillosetosum, G. glabriusculum, and G. sungpanense. They appear well separable from G. acutum.

Galium acutum and G. rebae are very closely related taxa. Mill (loc. cit. 1996: 199) presented a differential table which has been incor-

porated into the present descriptions. Nevertheless, from the material available now, it appears that only flower color (white or greenish white in G. acutum and reddish crimson in G. rebae) is really decisive for their separation. Furthermore, at lower elevations, G. acutum appears linked to G. asperifolium var. sikkimense. Cufodontis (Oesterr. Bot. Z. 89: 239. 1940) has described such transitional forms with longer and ± retrorsely aculeolate stems and hairy corolla lobes from the Indian Himalaya as G. acutum var. trichanthum Cufodontis.

Mill (loc. cit. 1996: 194–198) considered Galium acutum to be restricted to the NW Himalaya except for one provisionally identified specimen from Sikkim. In contrast, specimens studied by us from the herbaria PE, KUN, and WU clearly show that G. acutum extends much further to the east, reaching Yunnan and Sichuan.

Galium himalayense was regarded as a synonym of G. acutum by Cufodontis (loc. cit.: 239–243). Mill (loc. cit. 1996: 195; loc. cit. 1999: 831–832) agreed but maintained the taxon as G. acutum var. himalay-ense and described its sympatric occurrence with G. acutum var. acutum throughout the NW Himalaya. As Mill did not consider the possible occurrence of flower dimorphism in G. acutum, it remains uncertain whether the flower and stigma size differences listed are possibly cor-related with male and female plants or simply correspond to hermaph-roditic variants within the morphological amplitude of the species. In order to stimulate such studies and to clarify the distribution of the two taxa in China, a key and descriptions (according to Mill, loc. cit. 1996) follow:

1a. Open corollas 2.3–3.5 mm in diam.; stigmas united to near middle, in total length subequal to stamens ......................... 1a. var. acutum

1b. Open corollas 1.2–2.3 mm in diam.; stigmas united only shortly at base, in total length shorter than stamens ....... 1b. var. himalayense

1a. Galium acutum var. acutum

尖瓣拉拉藤(原变种) jian ban la la teng (yuan bian zhong)

Galium asperifolium Wallich var. setosum Cufodontis.

Leaves of main stems with blades mucronate at apex, mucro 0.2–0.45 mm. Corolla 2.3–3.5 mm in diam.; lobes 1.9–4 × as long as wide, mucronate with mucro 0.15–0.3 mm. Stig-mas united to near middle, in total length subequal to stamens. Fl. and fr. Jul–Oct.

Fl. China 19: 104–141. 2011.

Mountain rocks and slopes; 2000–4100 m. ?Sichuan, Xizang, ?Yunnan [India, Nepal, Pakistan].

1b. Galium acutum var. himalayense (Klotzsch & Garcke) R. R. Mill, Edinburgh J. Bot. 53: 195. 1996.

喜玛拉雅尖瓣拉拉藤 xi ma la ya jian ban la la teng

Galium himalayense Klotzsch & Garcke, Bot. Ergebn. Reise Waldemar, 88. 1862.

Leaves of main stems with blade submucronate at apex, mucro 0.15–0.2 mm. Corolla 1.2–2.3 mm in diam.; lobes 1.4–2.75 × as long as wide, acute or submucronate with mucro to 0.1 mm. Stigmas united only shortly at base, in total length shorter than stamens. Fl. and fr. Jul–Oct.

Mountain rocks and slopes; 2000–4100 m. ?Sichuan, Xizang, ?Yunnan [India, Nepal].

2. Galium aparine Linnaeus, Sp. Pl. 1: 108. 1753, s.s.

原拉拉藤 yuan la la teng

Herbs, annual, procumbent or clambering. Stems 30–90 cm high, 4-angled, 1–4 mm in diam., branched from base, retrorsely aculeate along angles, glabrescent to pilose at nodes. Leaves at middle stem region in whorls of 6–10, subsessile; blade drying papery, narrowly oblanceolate to narrowly oblong-oblanceolate, 10–60 × 3–10 mm, usually somewhat pilosulous or hispidulous adaxially, retrorsely aculeolate along midrib abaxially, base acute, margins flat to thinly revolute, retrorsely aculeolate, apex acute and shortly mucronate; vein 1. Inflores-cences terminal and axillary, cymes 2- to several flowered; axes glabrous to aculeolate; bracts ± leaflike or none, 1–5 mm; peduncles 1–5 cm; pedicels 1–30 mm, finally elongating and sometimes curved directly under fruit. Ovary subglobose, 0.3–0.5 mm, with uncinate trichomes. Flowers hermaphroditic. Co-rolla yellowish green or white, rotate, 1.5–2 mm in diam.; lobes 4, triangular to ovate, acute. Mericarps subglobose to kidney-shaped, 2.5–5 mm, with a dense cover of uncinate trichomes 0.4–1.2 mm from swollen base. Fl. Mar–Jul, fr. Apr–Nov.

Forest margins, riversides, meadows, open fields, farmlands; near sea level to 2500 m. Evidently rare in China and possibly only intro-duced [originally in W Eurasia and the Mediterranean, but today nearly worldwide as an adventive].

The Galium aparine group (G. sect. Aparine, formally part of G. sect. Kolgyda s.l.) forms an annual, extremely polymorphic, and pre-dominantly autogamous polyploid complex, also called G. aparine s.l. or G. aparine agg. One has to consider as possible perennial ancestors the morphologically very close E Asiatic taxa (e.g., G. sungpanense: see there) and other annuals, such as the Aegean endemic G. monachinii Boissier & Heldreich (2x, 2n = 22) and the Eurasian and African G. spurium (2x and 4x, 2n = 20, 40). By allopolyploidy they apparently have contributed in the Mediterranean and W Eurasia to G. aparine s.s. (4x, 6x, and 8x with ± euploid and slightly oscillating aneuploid chromosome numbers), which today has become a nearly worldwide weed (Ehrendorfer et al., Fl. Iranica 176: 239. 2005).

Many authorities, including Cufodontis (Oesterr. Bot. Z. 89: 245–247. 1940) and W. C. Chen (in FRPS 71(2): 234–237. 1999), have treated all these plants under Galium aparine s.l. and recognized four varieties: var. aparine, var. echinospermum, var. leiospermum, and var. tenerum. Whereas the first refers to G. aparine s.s. described above, the

latter three should be assigned to G. spurium (see there). Here, we follow the narrow circumscription of G. aparine s.s. and the specific separation of G. spurium outlined above, in spite of occasional diffi-culties in separating the two taxa on the basis of flower and mericarp size. A relevant survey of extensive Chinese material at the herbaria PE and KUN has clearly shown the common and wide occurrence of G. spurium in comparison with the rare and only occasional documentation of G. aparine s.s. Only future karyosystematic studies on the group in E Asia will clarify their distribution and ecological position. With respect to the common confusion of members of the G. aparine group with other annual and perennial taxa of Galium see G. spurium.

3. Galium asperifolium Wallich in Roxburgh, Fl. Ind. 1: 381. 1820.

楔叶律 xie ye lü

?Galium cavaleriei H. Léveillé; ?G. esquirolii H. Léveillé.

Herbs, perennial, weak to climbing or trailing, usually much branched. Stems 20–70 cm, 4-angled to 4-winged, vil-losulous to hirtellous and/or sparsely aculeolate to smooth. Leaves on main stems in whorls of up to 6(–8), sessile or with very short (ca. 1 mm) petiole; blade drying papery to leathery, adaxially dark green and shiny, abaxially paler, oblanceolate-oblong, oblanceolate, or obovate, (5–)10–20(–25) × (1–)1.5–4(–6) mm, adaxially scaberulous, hirtellous to glabrous, abaxi-ally densely villosulous, hirsute, pilose to glabrous, base acute to cuneate, margins retrorsely aculeolate and ± hairy, flat to thinly revolute, apex obtuse, rounded, truncate, or emarginate and shortly mucronate; vein 1. Inflorescences ± paniculate, up to 18 cm, expanding through growing season, with terminal and axillary, several- to many-flowered cymes; peduncles glabrous to rarely villosulous, regularly spreading to divaricate, with a dichasial branching pattern, at most nodes with leaflike bracts (1–4 mm); pedicels 0.2–2.5 mm. Ovary obovoid, 0.2–0.3 mm, mostly glabrous or smooth, but sometimes also verrucose, hir-tellous, or with undeveloped uncinate trichomes. Corolla greenish white or yellow, rotate, 1.5–2 mm in diam., glabrous, lobed for 2/3 or more; lobes 4, triangular-ovate, filamentous-aristate (rarely only acute). Mericarps ellipsoid, 1–2 mm, gla-brous and smooth or rarely granular-tuberculate, hirtellous, or with appressed to spreading hooked trichomes, on pedicels often slightly elongating to 4 mm. Fl. and fr. (May–)Jun–Sep(–Oct).

Mountain slopes, farmland sides, riversides and beaches, grass-lands, forests, thickets, ditch sides, open fields, meadows; 400–3500 m. Guangxi, Guizhou, Hubei, Hunan, Sichuan, Xizang, Yunnan [Afghani-stan, Bangladesh, Bhutan, India, Nepal, Pakistan, Sri Lanka, Thailand].

Galium asperifolium is an exceedingly variable and widespread species and was the first described from a larger assembly of taxa, here called G. asperifolium group and provisionally placed into G. sect. Tra-chygalium (but certainly not into G. sect. Leptogalium as in Yamazaki, Fl. Japan 3a: 238–239. 1993, or into G. sect. Leiogalium as in W. C. Chen, FRPS 71(2): 271. 1999). According to Cufodontis (Oesterr. Bot. Z. 211–251. 1940), Nazimuddin and Ehrendorfer (Pl. Syst. Evol. 155: 71–75. 1987), Mill (Edinburgh J. Bot. 53: 193–213. 1996; Fl. Bhutan 2(2): 825–834. 1999), Ehrendorfer et al. (Fl. Iranica 176: 194. 2005), and the present treatment, the center of diversity of the G. asperifolium group lies in the E Himalaya and SW China. Within this area, two sub-groups of taxa can be recognized, one with larger plants, longer than 10 mm middle stem leaves, and many-flowered cymes from lower eleva-

Fl. China 19: 104–141. 2011.

tions (1), the other with more condensed growth, shorter middle stem leaves, and 1- to few-flowered cymes from higher elevations (2). Both subgroups include taxa with whitish, yellowish to greenish, and others with reddish, purplish, or brownish flower color. The (sub)alpine sub-group (2) is briefly surveyed under G. acutum. Subgroup (1), discussed here, is represented by taxa with ± whitish flowers: G. subfalcatum Na-zimuddin & Ehrendorfer and G. campylotrichum Nazimuddin & Ehren-dorfer in the W Himalaya of Pakistan and the widespread G. asperi-folium with its var. sikkimense (= G. sikkimense) in the C and E Hima-laya (including Bhutan). In addition, subgroup (1) includes taxa with a more easterly distribution, extending from China into the remaining parts of E Asia (including Japan): G. dahuricum (including G. comarii, G. manshuricum, G. niewerthii, and G. pseudoasprellum), G. prattii, G. taiwanense, and G. tokyoense. Subgroup (1) taxa with reddish flowers are G. blinii in SW China and the newly described Bhutan endemic G. craticulatum R. R. Mill (see also Mill, loc. cit. 1996; loc. cit. 1999).

Because of its great variability with respect to habit and the in-dumentum of stems, leaves, and fruit, Galium asperifolium is often not easily separable from its closest relatives, and transitional forms occur. Its best differential characters are the many-flowered, divaricate, dist-ally dichasial branching, and strongly bracteate cymes and the small yellowish-greenish flowers with aristate corolla lobes. Galium blinii mainly deviates by larger, reddish flowers and non-aristate corolla lobes. The filiform and flexuose peduncles and pedicels separate G. dahuricum, and the less bracteate inflorescences and larger flowers separate G. prattii, G. taiwanense, and G. tokyoense. To the taxa of the (sub)alpine subgroup (2, e.g., G. acutum) G. asperifolium is linked particularly through its var. sikkimense.

In addition to its natural complexity, the taxonomy of the Galium asperifolium group is rendered difficult by a number of badly described and insufficiently documented species created by H. Léveillé. On the basis of the studies by Cufodontis (loc. cit.), Lauener and Ferguson (Notes Roy. Bot. Gard. Edinburgh 32: 103–115. 1973), Mill (loc. cit. 1996), and our own judgment, we suggest to dispose of them in the following way: Galium blinii is maintained as a separate species (pos-sibly with G. bodinieri and G. quinatum as synonyms), whereas G. cavaleriei and G. esquirolii (and “G. cuneatum H. Léveillé,” though a nomen nudum) are provisionally assigned as synonyms to G. asperi-folium s.l. (including var. sikkimense); G. comarii and G. niewerthii are treated as synonyms of G. dahuricum.

The following schematic key to the varieties of Galium asperi-folium corresponds to W. C. Chen (loc. cit.: 271–274) who mainly fol-lowed Cufodontis (loc. cit.: 239–240). Only G. asperifolium var. seto-sum has been eliminated because it clearly is a synonym of G. acutum. Individuals with uncinate trichomes on ovaries and fruit, but otherwise identical to typical G. asperifolium, are reported here for the first time. They still lack a varietal name and are provisionally accommodated under var. asperifolium. In contrast to Mill (loc. cit. 1996; loc. cit. 1999), G. sikkimense is here again reduced to varietal status under G. asperifolium, following Cufodontis (loc. cit.: 241). Forms of this species with ± glabrescent stems dominate to the east of its wide distribution area, but intraspecific and local variation of stem indumentum is so ex-tensive and continuous as to make this character useless as a basis for specific separation. Mill (loc. cit. 1996: 201–212) assumed G. asperi-folium var. asperifolium to be most common in C and W Nepal and to be replaced by var. sikkimense toward the east. This statement is in con-flict with the data on distribution in China from Chen (loc. cit.) and our own observations presented below. In view of the taxonomical com-plexity of the G. asperifolium group and the common misinterpretation of its members, further careful studies are obligatory.

1a. Plants stout, often clambering; stems usually villosulous to hirtellous and ± densely

retrorsely aculeolate; leaf blade relatively large, often oblanceolate-oblong, ± hairy and marginally retrorsely aculeolate; inflorescence large, many flowered; corolla lobes aristate. 2a. Ovary and fruit glabrous, smooth,

granular-tuberculate, or with uncinate trichomes .................... 3a. var. asperifolium

2b. Ovary and fruit hirtellous .......... 3b. var. lasiocarpum 1b. Plants often ± smaller; stems with ±

reduced indumentum; leaf blade smaller, often more lanceolate, more weakly retrorsely aculeolate to smooth along margins; inflorescence with fewer flowers; corolla lobes apiculate to acute. 3a. Mericarps smooth ......................... 3c. var. sikkimense 3b. Mericarps granular-tuberculate

................................................ 3d. var. verrucifructum

3a. Galium asperifolium var. asperifolium

楔叶律(原变种) xie ye lü (yuan bian zhong)

Galium pseudohirtiflorum H. Li.

Plants stout, often clambering. Stems 20–70 cm, much branched, villosulous or hirtellous, ± retrorsely aculeolate. Leaf blade often larger, oblanceolate-oblong, mostly ± hirsute and with margins retrorsely aculeolate. Inflorescences large, with many-flowered cymes. Corolla lobes filamentous-aristate. Ovary and fruit glabrous and smooth, verrucose, or rarely with uncinate trichomes. Fl. and fr. Jun–Sep.

Mountain slopes; 1200–3000 m. Guizhou, Sichuan, Xizang, Yun-nan [Afghanistan, Bangladesh, India, Nepal, Pakistan, Sri Lanka, Thai-land].

3b. Galium asperifolium var. lasiocarpum W. C. Chen, Acta Phytotax. Sin. 28: 303. 1990.

毛果楔叶律 mao guo xie ye lü

Ovary and fruit hirtellous. Fl. and fr. May–Oct.

● Mountain slopes, farmland sides, riversides, forests; 1400–3200 m. Guangxi, Guizhou, Yunnan.

Mill (loc. cit. 1996) commented on the extensive indument vari-ation on all organs of Galium asperifolium throughout most of its range but did not mention any occurrence of straight hairs on fruit in this species group, nor has anyone else besides W. C. Chen. Therefore, this variety is only provisionally included here. Its densely hirsute ovary and fruit suggests that it may belong to another species, possibly G. pusillo-setosum.

3c. Galium asperifolium var. sikkimense (Gandoger) Cufo-dontis, Oesterr. Bot. Z. 89: 241. 1940.

小叶律 xiao ye lü

Galium sikkimense Gandoger, Bull. Soc. Bot. France 66: 307. 1920.

Herbs, perennial, weak to clambering or trailing. Stems 20–60 cm, usually much branched, sparsely hairy and retrorsely aculeolate to smooth. Leaves often smaller and narrower, less

Fl. China 19: 104–141. 2011.

hairy and retrorsely aculeate to ± glabrous and smooth. Inflo-rescences large to medium sized with many- to several-flow-ered cymes. Corolla lobes apiculate to acute. Ovary and fruit glabrous and smooth. Fl. Jun–Sep, fr. Jul–Oct.

Mountain slopes, river beaches, ditch sides, open fields, grass-lands, meadows, thickets, forests; 400–3200 m. Guangxi, Guizhou, Hu-bei, Hunan, Sichuan, Xizang, Yunnan [Bhutan, India, Nepal].

3d. Galium asperifolium var. verrucifructum Cufodontis, Oesterr. Bot. Z. 89: 241. 1940.

滇小叶律 dian xiao ye lü

Similar to Galium asperifolium var. sikkimense but with granular-tuberculate fruit. Fl. and fr. Aug–Oct.

● Mountain slopes, grasslands, thickets; 2300–3500 m. Sichuan, Xizang, NW Yunnan.

The type material of this variety from Yunnan (particularly Han-del-Mazzetti 9697, WU) consists of transitional forms toward Galium acutum.

4. Galium asperuloides Edgeworth, Trans. Linn. Soc. London 20: 61. 1846.

车叶律 che ye lü

Herbs, perennial, emerging from filiform reddish rhi-zomes. Stems weak but generally erect, 10–45 cm tall, 4-an-gled, glabrous and smooth, except hispidulous at nodes. Middle stem leaves and leaflike stipules in whorls of (6 or)7 or 8, sub-sessile to very shortly petiolate; blade drying papery or mem-branous, remaining green, elliptic to narrowly oblong-oblanceo-late or lanceolate, (10–)20–50(–60) × 3–13 mm, length/breadth index mostly 3.5–4.5, glabrescent, base acute or cuneate, midrib smooth or rarely retrorsely aculeolate, margins and upper leaf side with antrorse microhairs, apex obtuse or rounded and abruptly apiculate; vein 1. Inflorescences terminal and some-times in axils of upper leaves with few- to several-flowered cymes; axes glabrous, smooth; bracts none or few, 1–2 mm; pedicels 0.5–5 mm. Ovary ovoid, 0.5–0.8 mm, with uncinate trichomes. Corolla white or light greenish, rotate, 2.5–3.8 mm in diam., lobed for 3/4 or more; lobes 4, ovate, acute. Mericarps ellipsoid, 1.8–2.5 mm, with dense uncinate trichomes 0.6–0.8 mm, on fruiting pedicels elongating to 10 mm. Fl. Apr–Aug, fr. May–Sep.

Forests on mountain slopes, thickets, ditch sides, along rivers, meadows; 1500–2800 m. Expected in Xizang [Afghanistan, India, Kashmir, Pakistan].

Galium asperuloides was previously circumscribed more broadly to include as subspecies plants treated here as G. hoffmeisteri. The speci-fic status of the latter is well justified (Ehrendorfer et al., Fl. Iranica 176: 193–194. 2005; see comments and differential characters under that species). When the two taxa are classified as one species, the “typical” plants have to be called G. asperuloides subsp. asperuloides. Vegetative plants are very similar to G. odoratum. Together with G. echinocarpum from Taiwan and others they constitute G. sect. Hylaea.

5. Galium baldensiforme Handel-Mazzetti, Symb. Sin. 7: 1029. 1936.

玉龙拉拉藤 yu long la la teng

Herbs, perennial, tender, caespitose. Stems ascending, 3–12(–25) cm tall, 4-angled, glabrous and smooth or rarely slightly retrorsely aculeolate. Leaves in whorls of up to 5 or 6, sessile; blade drying papery and blackish, oblanceolate to nar-rowly obovate, 2–7 × 1–3 mm, mostly glabrous and smooth, but occasionally with straight hairs abaxially or marginally slightly retrorsely aculeolate, base cuneate, margin flat, apex acute and often shortly mucronate; vein 1. Inflorescences with cymes terminal and in axils of upper leaves, 1- or usually 2- or 3-flowered; peduncles up to 18 mm and pedicels 2–5 mm, gla-brous and smooth, elongating during fruit development. Ovary subglobose, ca. 0.7 mm, densely covered by undeveloped tri-chomes. Corolla pale green, rotate, ca. 2 mm in diam., glabrous; lobes 4, triangular, acute. Mericarps ellipsoid, ca. 2 mm, with dense brownish yellow uncinate trichomes ca. 0.7 mm. Fl. Aug, fr. Aug–Oct.

● Rocky slopes, meadows, frost heave sites, river floodplains in mountains; 2800–4300 m. Qinghai, Sichuan, Xizang, Yunnan (Lijiang).

Galium baldensiforme belongs to the alpine subgroup (2) of the G. asperifolium complex within G. sect. Trachygalium s.l. (see under G. acutum and G. asperifolium). It has been widely misidentified with re-lated taxa, e.g., with G. glabriusculum from which it differs in its more oblanceolate, thinner leaves, blackening when dried, and a slight ten-dency toward more indumentum.

Two collections from Xizang (H. Li 1978-07-22 and Y. T. Chang & Lang, Nie-La-Mu, 1966-06-25, both from PE) differ from typical Galium baldensiforme by purple flowers (reminiscent of G. rebae), ± straight whitish hairs on its (still young) ovaries, and scattered straight hairs on the upper and lower leaf sides (mainly midvein, but glabrous along leaf margins). After closer inspection and field studies, these pop-ulations may very well deserve species rank.

A specimen from Sichuan (Dege Co., anonymous collector 7029, PE) with stronger indumentum, subleathery, broadly lanceolate leaves with retrorsely aculeolate margins, and aristate corolla lobes apparently links Galium baldensiforme with G. asperifolium var. sikkimense.

6. Galium blinii H. Léveillé, Bull. Acad. Int. Géogr. Bot. 25: 48. 1915.

五叶拉拉藤 wu ye la la teng ?Galium bodinieri H. Léveillé; ?G. quinatum H. Léveillé

& Vaniot.

Herbs, perennial, weak to climbing, trailing, or matted. Stems usually much branched, 20–70 cm, 4-angled, retrorsely aculeolate to glabrescent. Middle stem leaves in whorls of 6–8, subsessile; blade drying papery or leathery, often blackening, linear-oblong to broadly (ob)lanceolate, (5–)10–22(–30) × (1–)2–4.5(–5.5) mm, adaxially and particularly abaxially ± rough, base acute to cuneate, margin flat to thinly revolute, densely retrorsely aculeolate, ± gradually narrowed into acute apex; vein 1. Inflorescences with terminal and axillary, several-flowered cymes 2–5 cm; axes ± glabrous, often slightly di-varicate, with small bracts on lower branches only; pedicels (0.2–)1–3(–5) mm. Ovary obovoid, 0.2–0.3 mm, glabrous or with undeveloped trichomes. Corolla red to purple or violet (only very rarely white), rotate, 1.5–2.5(–3) mm in diam., gla-brous; lobes 4, triangular-ovate, acute. Mericarps ovoid, 1–2 mm, glabrous, smooth to verrucose, or sometimes with ap-pressed or spreading uncinate trichomes. Fl. Jun–Sep, fr. Jul–Oct.

Fl. China 19: 104–141. 2011.

● Mountain slopes, river beaches, ditch sides, open fields, grass-lands, meadows, thickets, forests; 800–3000 m. Guizhou, Hubei, Shaan-xi, Sichuan, Xizang, Yunnan.

Galium blinii is a critical taxon that belongs to the lower elevation subgroup (1) of the G. asperifolium group (see there). It was considered a synonym of G. asperifolium var. sikkimense by Cufodontis (Oesterr. Bot. Z. 89: 241. 1940), Lauener and Ferguson (Notes Roy. Bot. Gard. Edinburgh 32: 106. 1973), and W. C. Chen (in FRPS 71(2): 273. 1999). Only Mill (Edinburgh J. Bot. 53: 204. 1996) commented on its reddish purple flowers and other differential characters, regarded it as a distinct species, and proposed to use its old but so far neglected name. Up to now, specimens of this rather widespread taxon were named as G. asperifolium var. sikkimense, G. pseudoasprellum var. densiflorum, etc. Analyses of a considerable number of relevant specimens from PE, KUN, and WU support Mill’s interpretation and led to the above, more elaborate description. It shows that G. blinii is quite variable with re-spect to leaf shape and ovary/fruit indumentum but relatively well char-acterized not only by its reddish purple flowers but also by height, leaf size, retrorsely aculeolate stems and leaf margins, and the usually medi-um-sized and only small-bracteate cymes, relatively short and firm post-floral peduncles and pedicels, and larger flowers with acute (but not aristate) lobes. This allows separation from its closest relatives, G. prattii and G. asperifolium (where transitional forms occur), but also from G. dahuricum and G. tokyoense.

Further studies will have to show to what extent the following, also reddish purplish flowering taxa from SW China can be separated from Galium blinii: G. craticulatum was described as an endemic from the high mountains of Bhutan (Mill, loc. cit.: 202) and said to differ from G. blinii in its present circumscription by completely glabrous stems with conspicuous, vein-marked wings, less retrorsely aculeolate leaves, larger flowers, and longer filaments; G. bodinieri, also with red-dish flowers, according to Mill (loc. cit.: 204–205) is reminiscent of G. craticulatum (particularly in its broadly winged stems) and may represent a link between G. blinii and the alpine, more condensed and shorter leaved G. rebae; the purple-flowered G. quinatum (not men-tioned by Mill) was very poorly described and is listed above as a pos-sible synonym of G. blinii, but types have neither been seen by Lauener and Ferguson (loc. cit.: 107) nor by us.

7. Galium boreale Linnaeus, Sp. Pl. 1: 108. 1753, s.l.

北方拉拉藤 bei fang la la teng

Herbs, perennial, erect, 20–65 cm tall. Stems 4-angled, usuually glabrous, rarely shortly hairy, hispidulous at nodes, angles thickened. Leaves in whorls of 4, sessile or subsessile; blade drying papery or thinly leathery, linear-lanceolate or lan-ceolate to ovate, (10–)15–40(–80) × (1–)3–15 mm, glabrous or sparsely puberulent to hispidulous and/or pilose, abaxially never with striate to punctate glandular idioblasts, base cuneate to sub-rounded, margins usually revolute and antrorsely scaberulous to hispidulous, apex acute or usually narrowly tapered then obtuse to rounded at very tip; principal veins palmate, 3. Inflores-cences terminal, elongate or broadly paniculiform, 2–15 cm, with several- to many-flowered cymes in axils of uppermost leaves and terminal; peduncles glabrous or puberulent at nodes, smooth or scaberulous; bracts ligulate, lanceolate, or elliptic, 1–4 mm; pedicels 0.5–2 mm elongating in fruit to 3.5 mm. Ovary subglobose, 0.8–1 mm, glabrous or sparsely to densely strigil-lose to pilosulous. Corolla white or pale yellow, rotate, 3–4 mm in diam., glabrescent, lobed for 3/4 or more; lobes 4, ovate-lan-ceolate, acute. Mericarps subglobose, 1–2 mm, pericarp firmly attached but sometimes ± inflated, glabrous or ± densely hairy

with ± appressed, ascending, or spreading, straight or curved, but hardly truly uncinate trichomes 0.3–0.5 mm. Fl. May–Aug(–Sep), fr. (May–)Jun–Oct.

Open forests and thickets, mountain slopes, grasslands, meadows, open fields, ditch sides, river valleys and beaches, swamps, farmland sides, wastelands; 200–4600 m. Gansu, Hebei, Heilongjiang, Henan, Jilin, Liaoning, Nei Mongol, Ningxia, Qinghai, Shaanxi, Shandong, Shanxi, Sichuan, Xinjiang, Xizang, Yunnan [Afghanistan, India, Japan, Kashmir, Korea, Mongolia, Pakistan, Russia; SW Asia (Armenia, Iran), Europe, North America].

The name Galium boreale, as used here in a wide sense, cor-responds to a widespread and polymorphic, still insufficiently studied N Hemisphere polyploid complex (Ehrendorfer et al., Fl. Iranica 176: 179–181. 2005) within G. sect. Platygalium s.l. In China, another species of this section with much smaller flowers, G. kinuta, can be separated from this G. boreale aggregate only with difficulties, because the two are linked by intermediate (and possibly hybrid) populations (see under G. kinuta).

Within the Galium boreale aggregate and the flora of China, W. C. Chen (in FRPS 71(2): 260–263, 285. 1999) recognized only G. boreale Linnaeus s.l. with numerous infraspecific taxa and G. turkestanicum, whereas 11 species in three series were listed for the flora of the former Soviet Union by Pobedimova et al. (Fl. URSS 23: 345–354. 1958). From these only G. turkestanicum is fully accepted here (G. ussuriense and G. rubioides are cited as synonyms under G. boreale var. lanceo-latum and G. boreale var. rubioides). Furthermore, and according to Pobedimova et al. (loc. cit.), G. amblyophyllum Schrenk, G. amurense Pobedimova, and G. septentrionale Roemer & Schultes can be expected to occur in China. With the exception of the briefly mentioned G. sep-tentrionale, they were not considered by W. C. Chen in FRPS and are only mentioned here. As a competent treatment of the G. boreale aggre-gate is not yet possible, we follow the schematic taxonomic differenti-ation proposed by Cufodontis (Oesterr. Bot. Z. 89: 225–228. 1940) and accepted by W. C. Chen (loc. cit.). This scheme defines numerous varieties according to leaf shape and the density, type, and distribution of indumentum on leaves, ovaries, and fruit. These varieties form a morphologically ± continuous series, linking the extremes: G. boreale var. rubioides with large ovate leaves and a broadly paniculate inflores-cence and G. boreale var. intermedium with much smaller lanceolate leaves and an elongated narrow inflorescence. The following key and short descriptions are presented here for reference, to facilitate compari-son, and to stimulate future studies.

1a. Ovary and fruit glabrous. 2a. Leaf blade pilose or scabrous

abaxially at least along veins. 3a. Leaf blade linear-lanceolate

or narrowly lanceolate ...... 7h. var. lancilimbum 3b. Leaf blade ovate-lanceolate

or ovate ................................... 7k. var. rubioides 2b. Leaf blade glabrous abaxially.

4a. Leaf blade linear-lanceolate or narrowly lanceolate .... 7d. var. hyssopifolium

4b. Leaf blade broadly lanceolate or ovate-lanceolate ................. 7g. var. lanceolatum

1b. Ovary and fruit ± hairy. 5a. Ovary and fruit sparsely hirtellous

or scabrous. 6a. Leaf blade linear-lanceolate

or narrowly lanceolate ...... 7e. var. intermedium 6b. Leaf blade broadly lanceolate

or ovate-lanceolate ....... 7j. var. pseudorubioides

Fl. China 19: 104–141. 2011.

5b. Ovary and fruit densely hirsute or tomentose. 7a. Leaf blade sparsely pubescent

or scabrous at least along veins abaxially. 8a. Leaf blade linear-lanceolate

or narrowly lanceolate ..... 7c. var. ciliatum 8b. Leaf blade broadly

lanceolate or ovate-lanceolate ..... 7f. var. kamtschaticum

7b. Leaf blade glabrous abaxially. 9a. Leaf blade less than

4 mm wide ......................... 7b. var. boreale 9b. Leaf blade 4–15 mm wide.

10a. Leaf blade 4–6 mm wide ......... 7a. var. angustifolium

10b. Leaf blade wider than 6 mm .............. 7i. var. latifolium

7a. Galium boreale var. angustifolium (Freyn) Cufodontis, Oesterr. Bot. Z. 89: 226. 1940.

狭叶砧草 xia ye zhen cao

Galium rubioides Linnaeus var. angustifolium Freyn, Oesterr. Bot. Z. 45: 341. 1895.

Leaf blade narrowly lanceolate, 4–6 mm wide, abaxially glabrous. Ovary and mericarps densely hirsute or tomentose. Fl. Jun–Aug, fr. Jul–Sep.

Mountain slopes, river valleys, swamps, grasslands, meadows; 500–3900 m. Hebei, Heilongjiang, Nei Mongol, Sichuan, Xinjiang, Xizang [Japan, Kashmir, Russia; NE Europe].

7b. Galium boreale var. boreale

北方拉拉藤(原变种) bei fang la la teng (yuan bian zhong)

Galium boreale var. vulgare Turczaninow.

Leaf blade narrowly lanceolate or linear-lanceolate, 1–3.9 mm wide, glabrous. Ovary and mericarps densely hirsute or tomentose with slightly curved, white trichomes. Fl. May–Aug, fr. Jun–Oct.

Forests, thickets, or tussocks on mountain slopes, ditch sides, grasslands, meadows; 700–3900 m. Gansu, Hebei, Heilongjiang, Jilin, Liaoning, Nei Mongol, Qinghai, Shandong, Shanxi, Sichuan, Xinjiang, Xizang [India, Korea, Pakistan, Russia; Europe, North America].

7c. Galium boreale var. ciliatum Nakai, J. Jap. Bot. 15: 340. 1939.

硬毛拉拉藤 ying mao la la teng Leaf blade linear-lanceolate or narrowly lanceolate, 1–6

mm wide, abaxially sparsely hairy or scabrous at least along veins. Ovary and mericarps densely hirsute or tomentose. Fl. Jun–Sep, fr. Jul–Oct.

Mountain slopes, river beaches, ditch sides, open fields, meadows; 200–4600 m. Gansu, Hebei, Heilongjiang, Jilin, Liaoning, Nei Mongol, Ningxia, Qinghai, Shaanxi, Shanxi, Sichuan, Xinjiang, Xizang, Yunnan [Japan, Russia; Europe (Finland, Romania), North America].

7d. Galium boreale var. hyssopifolium (Hoffmann) Candolle, Prodr. 4: 600. 1830.

斐梭浦砧草 fei suo pu zhen cao

Galium hyssopifolium Hoffmann, Deutschl. Fl., Dritter Jahrgang, 71. 1800; G. boreale f. hyssopifolium (Hoffmann) B. Boivin; G. boreale subsp. hyssopifolium (Hoffmann) Schübler & G. Martens; G. rubioides var. hyssopifolium (Hoffmann) Per-soon.

Leaf blade linear-lanceolate or narrowly lanceolate, 1–6 mm wide, glabrous abaxially. Ovary and mericarps glabrous. Fl. and fr. Jun–Aug.

Mountain slopes, grasslands; 1800–2300 m. Sichuan, Xinjiang [Europe].

This variety probably has a wider geographic range.

7e. Galium boreale var. intermedium Candolle, Prodr. 4: 601. 1830.

新砧草 xin zhen cao Leaf blade linear-lanceolate or narrowly lanceolate, 1–6

mm wide, sparsely pubescent to glabrescent. Ovary and meri-carps sparsely hirtellous or scabrous. Fl. Jun–Jul, fr. Jul–Oct.

Mountain slopes, wastelands, forests, grasslands, meadows; 1500–1800 m. Gansu, Heilongjiang, Xinjiang [Russia; Europe].

7f. Galium boreale var. kamtschaticum (Maximowicz) Nakai, Bot. Mag. (Tokyo) 23: 103. 1909.

堪察加拉拉藤 kan cha jia la la teng Galium boreale f. kamtschaticum Maximowicz, Mém.

Acad. Imp. Sci. St.-Pétersbourg Divers Savans 9 [Prim. Fl. Amur.]: 141. 1859; G. boreale var. koreanum Nakai.

Leaf blade broadly lanceolate or ovate-lanceolate, 6–10 mm wide, abaxially sparsely hairy or scabrous at least along veins. Ovary and mericarps densely hirsute or tomentose. Fl. and fr. Jun–Sep.

Mountain slopes, farmland sides, riversides, river valleys, grass-lands, meadows; 800–2400 m. Heilongjiang, Henan, Jilin, Liaoning, Nei Mongol, Shaanxi, Shanxi, Sichuan, Xinjiang [Kashmir, Korea, Mongolia, Russia; C Asia (“Turkestan”), NE Europe].

Nakai (Bot. Mag. (Tokyo) 23: 103. 1909) commented on Japanese plants and presented a key to Galium, including “Galium boreale var. kamtschaticum Maximowicz” with no further information. Some au-thors have considered this a validly published combination (e.g., in Go-vaerts et al., World Checkl. Rubiaceae; http://www.kew.org/wcsp/rubia-ceae/; accessed on 16 Nov 2007), whereas others have not (e.g., W. C. Chen, loc. cit.: 263) and have instead cited J. Coll. Sci. Imp. Univ. Tokyo 31: 498. 1911 (sometimes called Fl. Koreana 2) as the place of publication. Nakai’s new co